Effect of group size and composition on individual behavior, group dynamics and population regulation in the Arabian babbler (Turdoides squamiceps) Oded Keynan A dissertation submitted to Macquarie University in application for the degree of Doctor of Philosophy. November 2014 Department of Biology 2 3 Table of contents Abstract .............................................................................................................................................. 6 Statement of candidate ....................................................................................................................... 7 Preface ................................................................................................................................................ 8 Acknowledgements .......................................................................................................................... 10 1. Introduction .............................................................................................................................. 15 1.1 Cooperative breeding ............................................................................................................ 16 1.2 Social foraging and social learning ...................................................................................... 17 1.3 Individual success, group dynamics and Allee effects ....................................................... 20 1.4 Thesis structure ..................................................................................................................... 23 2. General methods ...................................................................................................................... 25 2.1 Study species: ........................................................................................................................ 26 2.2. Study site ............................................................................................................................... 31 2.3. The Arabian babbler research station ............................................................................... 34 2.4 Data collection ....................................................................................................................... 35 2.5 Statistical analysis ................................................................................................................. 39 3. Social foraging strategies and acquisition of novel foraging skills in cooperatively- breeding Arabian babblers. ............................................................................................................................. 43 3.1 Abstract .................................................................................................................................. 44 3.2 Introduction ........................................................................................................................... 44 3.3. Materials & methods ........................................................................................................... 47 3.4. Results ................................................................................................................................... 54 3.5 Discussion .............................................................................................................................. 59 3.6 References .............................................................................................................................. 63 4 4. Task-dependent differences in innovation abilities by subordinate and dominant wild Arabian babblers ............................................................................................................................................ 69 4.1 Abstract ................................................................................................................................. 70 4.2 Introduction .......................................................................................................................... 70 4.3. Materials & methods ........................................................................................................... 74 4.4 Results ................................................................................................................................... 79 4.5. Discussion ............................................................................................................................. 84 4.6 References ............................................................................................................................. 89 5. Causes of variation in dominance acquisition, tenure, and lifetime reproductive success in a cooperatively breeding bird .............................................................................................................. 95 5.1 Abstract ................................................................................................................................. 96 5.2 Introduction .......................................................................................................................... 97 5.3 Materials & methods ............................................................................................................ 99 5.4. Results ................................................................................................................................ 107 5.5 Discussion ............................................................................................................................ 116 5.6 References ........................................................................................................................... 120 6. Long-term analysis of social and environmental influences on group dynamics in a cooperative breeder ............................................................................................................................................ 125 6.1 Abstract ............................................................................................................................... 126 6.2 Introduction ........................................................................................................................ 127 6.3 Materials & methods .......................................................................................................... 130 6.4 Results ................................................................................................................................. 138 1208.67 ........................................................................................ Error! 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Long-term population analysis reveals the presence of group and demographic Allee effects on a cooperatively breeding bird species. .............................................................................................. 159 5 7.1 Abstract ................................................................................................................................ 160 7.2 Introduction ......................................................................................................................... 161 7.3 Materials & methods .......................................................................................................... 163 7.4 Results .................................................................................................................................. 167 7.5 Discussion ............................................................................................................................ 172 7.6 References ............................................................................................................................ 174 8. General discussion ..................................................................................................................... 179 8.1 Overview .............................................................................................................................. 180 8.2 Social foraging, acquisition of novel skills and innovation .............................................. 181 8.3 The effect of social and environmental factors on individual fitness traits and group dynamics. ................................................................................................................................... 183 8.4 Inverse density dependence and Allee effects. .................................................................. 186 8.5 Conclusion ..........................................................................................................................